North American Fungi

Volume 5, Number 4, Pages 1-10 Published November 19, 2010

Aurantiopileus mayanensis a new and of polypore (, ) from Belize with connections to existing Asian species

James Ginns1, Daniel L. Lindner2, Timothy J. Baroni3, Leif Ryvarden4

11970 Sutherland Road, Penticton, BC V2A 8T8 Canada, 2 US Forest Service, Northern Research Station, Center for Forest Mycology Research, One Gifford Pinchot Dr., Madison, WI 53726-2398 USA,3 Department of Biological Sciences, P. O. Box 2000, State University of New York, College at Cortland, Cortland, NY 13045 USA,4 Department of Botany, University of Oslo, P. O. Box 1066, Blindern, NO-0316 Norway

Ginns, J., D. L. Lindner, T. J. Baroni, and L. Ryvarden. 2010. Aurantiopileus mayanensis a new genus and species of polypore (Polyporales, Basidiomycota) from Belize with connections to existing Asian species. North American Fungi 5(4): 1-10. doi: 10.2509/naf2010.005.004

Corresponding author: Timothy J. Baroni [email protected]. Accepted for publication September 17, 2010. http://pnwfungi.org Copyright © 2010 Pacific Northwest Fungi Project. All rights reserved.

Abstract: A new genus and species of polypore, Aurantiopileus mayanensis, is described from Central America based on a visually striking collection made in the cloud forests of the Maya Mountains in Belize. Phylogenetic analysis using nLSU sequences and distinctive morphological characters support the erection of a new genus to accommodate this species. Two Asian species formerly placed in are also considered to be members of this new genus and the new combinations are provided, along with a key to the three taxa now placed in Aurantiopileus.

Key words: Aurantiopileus mayanensis, Gloeoporus, Phlebia, brightly colored polypores, key to species of Aurantiopileus, Doyle’s Delight, Maya Mountains. 2 Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10

Introduction: During the second expedition to The following description of this new genus and the highest peak in the Maya Mountains of Belize species is accompanied by a key to species we (Doyle’s Delight), 20-30 new taxa of macrofungi include in this new genus. A full color image of were discovered. One of these collections is this new taxon is presented here and can also be presented here as a new genus and species of seen at: polypore based on morphological and molecular http://news.nationalgeographic.com/news/2008 evidence. The remote cloud forest habitat is not /10/081031-belize-mushroom-missions.html. easily accessible by normal forms of transportation (automobile/walking), but does Materials and Methods: The color codes lend itself to helicopter assisted drop-off cited are from Kornerup and Wanscher (1978). expeditions and two expeditions have now been The abbreviations for the herbaria where completed at this remote site (August 2004 and specimens are preserved follow Holmgren et al. August 2007). The remoteness of the area (1990). The standard mounting media for appears to play a role in the discovery of large examination of specimens of the Polyporaceae numbers of new and yet to be described taxa of and allied groups were used, i.e., Melzer's macrofungi. Reagent, 2% potassium hydroxide (KOH), and cotton blue in lactic acid. The formulae for these Baroni et al. (2007 and 2008) and Ryvarden et al. can be found in Kirk et al. (2001). Morphological (2009) have published on new taxa from this data are based upon examination of basidiomata remote site (one new genus and three new of Aurantiopileus mayanensis (T.J. Baroni species). Baroni et al. (2009) presented data on 10228), Gloeoporus dolosus Corner (E.J.H. three more new genera from Doyle’s Delight that Corner, 4.X66, E00159719) and G. pendens are now in preparation for publication. As we Corner (RSNB 5200, E00159705). and other experts in various groups sort through the 1,000 or so collections acquired during those Molecular data are based on basidiomata of two trips, more new taxa and reports will be Aurantiopileus mayanensis (T.J. Baroni 10228) forthcoming. and additional sequences from GenBank. Tissues for DNA extraction were fixed in the field using The Doyle’s Delight site is considered a lower fine mesh silica gel in Corning 2.0 ml Self montane (1100 meters amsl) subtropical wet Standing MicroCentrifuge tubes with o-ring seal forest (Holdridge et al. 1971). The forest is screw caps. DNA extraction from basidiomata composed of mixed tropical Angiosperm trees and PCR procedures followed Palmer et al. with some cloud forest elements rich with (2008). In brief, small fragments of basidiomata epiphytes. Some of the trees were Euterpe were excised with a sterile scalpel and placed in precatoria Mart., Colpothrinax cookii Read, 1.5 mL microcentrifuge tubes. DNA was Cyrilla racemiflora L., Sloanea floribunda extracted using GeneClean III kits (Qbiogene) Benth., Magnolia sp., Clusia sp., Neea sp., following manufacturer’s protocols. The internal Calatola sp., Quercus spp., etc. The downed and transcribed spacer (ITS) region of rDNA was PCR decaying decorticate log from which our sample amplified using the fungal-specific primer pair came was not readily identifiable. The rot caused ITS1F (Gardes and Bruns 1993) and ITS4 (White by Aurantiopileus mayanensis appeared to be a et al. 1990) and a section of the nuclear large white stringy rot typical of hardwood angiosperm subunit (nLSU) rDNA was amplified using substrates. primers LR16 and LR0R (Hopple and Vilgalys Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10 3

1994, Klonowska et al. 2003). PCR products were Maximum parsimony was implemented in PAUP purified and sequenced following Lindner and 4.0b10 (Swofford 2002). Heuristic searches were Banik (2009). Sequences were deposited in conducted with characters unordered and of GenBank with the following accession numbers: equal weight and gaps treated as missing data. HM772139 (partial nLSU) and HM772140 (ITS Default settings were used with the following region). exceptions: stepwise-addition option was set to random with 100 replicates, steepest descent was Sequences were aligned manually using Sequence used with the TBR branch swapping option and Alignment Editor (Se-Al) v2.0a9. The nLSU the number of trees saved was set to sequence obtained from Aurantiopileus automatically increase. Bootstrap support for mayanensis was aligned with 18 additional clades (Felsenstein 1985) was estimated from sequences from Genbank ( adusta 1000 heuristic searches with the same settings (Willd.) P. Karst. AF287848, Climacodon described above, with the exception that the septentrionalis (Fr.) P. Karst. AY684165, stepwise-addition option was set to random with Crustodontia chrysocreas (Berk. & M.A. Curtis) 50 replicates. Bayesian inference was Hjortstam & Ryvarden AY293199, Gloeoporus implemented in MrBayes 3.1.2 (Huelsenbeck and dichrous (Fr. ) Bres. EU118627, G. taxicola Ronquist 2001; Ronquist and Huelsenbeck 2003) (Pers.) Gilb. & Ryvarden AY586656, Laetiporus using default settings with the GTR model. One sulphureus (Bull.) Murrill EU402532, million generations were performed with samples Lilaceophlebia livida (Pers.) Spirin & Zmitr. taken in increments of 100. The first 2500 trees AF141624, L. subserialis (Bourdot & Galzin) (25%) were considered the “burnin” and were Spirin & Zmitr. AF141631, aurea (Fr.) excluded from construction of the consensus tree. J. Erikss. & Ryvarden AY586691, M. uda (Fr.) After running the analysis the standard deviation Donk AF141614, Phanerochaete chrysorhiza of the split frequencies was examined to confirm (Torr.) Budington & Gilb. AF139967, Phlebia it was below 0.01, the potential scale reduction acerina Peck AF141615, P. centrifuga P. Karst. factor was examined to confirm all parameters AF141618, P. radiata Fr. AF287885, P. rufa were close to 1.0, and the plot of the generations (Pers.) M.P. Christ. AF141628, P. tremellosa versus the log probability of the data (the log (Schrad.) Nakasone & Burds. AF141632, likelihood values) was examined to confirm it had Polyporus squamosus (Huds.) Fr. AF393069, reached the stationary phase. Scopuloides hydnoides (Cooke & Massee) Hjortstam & Ryvarden EU118665) and two Results sequences from Karl-Henrik Larsson (Mycoacia fuscoatra (Fr.) Donk and bispora Aurantiopileus Ginns, D. L. Lindner & T. J. (Stalpers) J. Erikss. & Ryvarden), indicated with Baroni, gen. nov. collection numbers beginning KHL or EL (pers. comm.). Sequences were selected for alignment Basidiomata comparate parva, quum recentia textura based on nLSU and ITS BLAST searches in succulenta vel gelatinoss, hymenophoro poroideo, GenBank, which indicated that contextu gelatinoso, cystidia tenui-tunicatis in trama and other members of the phlebioid clade within gelatinosa inclusis, systemate hyphali monomitico, the polyporoid clade (sensu Binder et al. 2005) fibulis praedita, basidiosporis parvis late ellipsoideis were the closest matches to Aurantiopileus (4-6 x 2.8 -4.6 um), inamyloideis; data molecularia, e mayanensis. Phylogenetic analyses were nLSU collecta, Aurantiopileum e generibus aliis performed only on nLSU data because ITS data proxime affinibus in clado phlebioideo inclusis could not be unambiguously aligned. separantia. 4 Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10

MycoBank#: MB 518879 Habitat: Associated with a white, stringy rot of a Typus: Aurantiopileus mayanensis Ginns, D. L. 1 ft (30 cm) diameter downed decorticated Lindner & T. J. Baroni hardwood log in mixed tropical cloud forest.

Basidiomata relatively small, with a Distribution: Belize, known only from the type fleshy/gelatinous texture when fresh, a poroid locality. hymenophore, gelatinous context and trama with thin walled cystidia embedded in the gelatinous Basidioma when fresh orange (7A-B7-8), soft, trama, hyphal system monomitic, with clamp gel-like, deeper orange over margin (8B8), connections, basidiospores small and broadly effused-reflexed or substipitate, 20-60 mm ellipsoid (4-6 x 2.8-4.6 m), inamyloid, and broad, circular when substipitate or dimidiate molecular data from nLSU separate when lacking a stipe. Pilei imbricate, confluent Aurantiopileus from other genera in the and attached together, intergrading. Pileus phlebioid clade with which it is most closely surface glabrous, rugulose, expallent hairs near allied. attachment white, slightly zonate. Pore surface orange; pores round or angular, very fine (2-4 per Aurantiopileus mayanensis Ginns, D. L. mm) and shallow (approximately 1 mm deep) or Lindner & T. J. Baroni sp. nov. (Figures 1-5) appearing somewhat rugulose-pitted or fluted. Pore edges fimbriate to dentate. Context Etymology: genus name after the orange pileus orange, watery, gel-like, 2 mm thick. and the species name commemorates the locality, Maya Mountains. When dried pileus surface white to pale brown, radially striate, striae fine, slightly glossy. Basidiocarpus sessilis, in vivo omnino carneus et Margin thin, hard, orange, plane or inrolled. aurantiacus. Pileus in sicco albus vel pallidus Context hard (horny), waxy, orange with pale brunneus, margo sterilis. Hymenophorus in brown, radiating striate, 1.0-2.5 mm thick. Pores sicco aurantiacus, poris, angularis, 2-4 per mm, orange, round to angular, (2-) 3 (-4) per mm. tubulis concoloribulus usque 1 mm long. Systema hypharum monomiticum, hyphis fibulatis, 3-6 Hyphal system monomitic. Hyphae with µm diam., tenu-tunicatis, indextrinoideis. clamp connections, walls hyaline, negative in Cystidius in trama, clavatus, 35-100 x 5-15 μm, Melzer’s, acyanophilous. Pore edges with hyphae paries tenuis, hyalinin, levibus. Basidiosporis in fascicles, 3.6-4.0 μm diameter, thin-walled. ellipsoideus vel latus ellipsoideus, (4.6-) 5.0-6.0 x Tramal and context hyphae with thickened, (3.4-) 3.6-4.2 (-4.6) μm, paries tenuis, hyalinin, gelatinous walls that merge into structureless levibus, inamyloides. mucilage and the hyphae appear to be imbedded in a gelatinous matrix. Pileus surface hyphae MycoBank #: MB 518880 sparse, in fascicles, forming a discontinuous, pallid layer composed of hyphae (3-) 4 (-5) μm Collection data: Belize: Cayo District: Maya diameter, walls thin to 0.5 μm thick. Mountains: Doyle's Delight Peak: North Ridge Gloeopleurous hyphae scattered in trama and Trail above Creek/Waterhole on north side, N 16 context, contents homogeneous, pale yellow in 30 14.3, W 89 03 04.3 (WGS84), elevation 1100 KOH, red in phloxine (unlike other hyphae), 3-6 meters, 28 Aug. 2007, coll. T. J. Baroni 10228 μm diameter, branched. Cystidia confined to (HOLOTYPE: CORT; ISOTYPES: BRH and tramal core, imbedded, typically in vertically CFMR). oriented fascicles, clavate, narrowly clavate or fusoid, 35-100 x 5-15 μm, walls thin, smooth, Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10 5 negative in Melzer’s, some with a thick, specimens of Aurantiopileus are a fleshy/ gelatinous coating, contents homogeneous, gelatinous texture, a poroid hymenophore and slightly refractive in KOH, dark blue in Cotton relatively small (less than 60 mm broad) Blue. Basidia clavate, 18-26 x 6-7 μm, four basidiomata. Microscopically the combination of sterigmate, each slender, 5 μm long. gelatinous context and trama, cystidia imbedded Basidiospores ellipsoid to broadly ellipsoid, in the trama, monomitic hyphal system, hyphae (4.6-) 5.0-6.0 x (3.4-) 3.6-4.2 (-4.6) μm (n = 20), with clamp connections, and small (4.0-6.0 x 2.8- walls smooth, thin, hyaline, negative in Melzer’s, 4.6 µm), mostly broadly ellipsoid basidiospores most containing one large oil drop. distinguish the genus. Molecular data from the nLSU of A. mayanensis (Fig. 6) also help to distinguish this genus, and indicate Aurantiopileus is distinct from other similar genera (e.g. Bjerkandera P. Karst., Gloeoporus Mont., Phlebia Fr.) and falls in the phlebioid clade within the polyporoid clade (sensu Binder et al. 2005).

Figure 5. Aurantiopileus mayanensis. T. J. Baroni 10228. Basidiomata, in situ. Figures 1-4. Aurantiopileus mayanensis. T. J. Baroni 10228 : Fig. 1. Basidiospores at 1000x, Similar embedded cystidia occur in several other DIC. Scale bar = 5 µm. Fig. 2. Sterile cells on species of the Polyporales. Two from the North pore edges at 400x, DIC. Scale bar = 10 µm. Fig. Temperate zone and microscopically similar to A. 3. Embedded tramal cystidia at 1000x, DIC. mayanensis are Phlebia radiata and P. rufa. In Scale bar = 20 µm. Fig. 4. Embedded tramal tropical Asia two species of Gloeoporus, in cystidia at 400x, DIC. Scale bar = 10 µm. addition to similar cystidia have partly gelatinized basidiomata and several microscopic features similar to those of A. mayanensis. These Discussion Asian species were described by Corner (1989) when he proposed thirteen new species of Morphological Considerations: The Gloeoporus from Malaysia and Japan. At the distinctive macroscopic features of fresh same time he transferred the type species of Bjerkandera and Skeletocutis Kotl. & Pouzar to 6 Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10

Gloeoporus. One result was the broadening of the dolosus are brownish black; a color that may be a circumscription of the genus Gloeoporus. result of their having been in alcohol-formalin. New combinations proposed: We prefer a Although the brownish black coloration is narrower generic concept and, based on their distinct from the pale orange brown basidiomata endotramal cystidia and predominately broadly of A. pendens, it may not be taxonomically ellipsoid basidiospores, transfer two species significant. We did not find any microscopic described in Gloeoporus by Corner (1989) to features to clearly distinguish A. dolosus from A. Aurantiopileus: pendens. The three Aurantiopileus species 1. Aurantiopileus dolosus (Corner) apparently occupy a similar niche (i.e., the Ginns & D. L. Lindner comb. nov. fragments of decomposed, hardly recognizable (Basionym: Gloeoporus dolosus Corner, 1989, wood and humus are attached to the Nova Hedwigia, Beihefte 96: 50) basidiomata). The basidiomata of A. mayanensis 2. Aurantiopileus pendens (Corner) shrunk somewhat on drying and become hard Ginns & D. L. Lindner comb. nov. and waxy but the margin and pores retained the (Basionym: Gloeoporus pendens Corner, 1989, distinctive orange color. Nova Hedwigia, Beihefte 96: 54). Key to the species of Aurantiopileus: Corner stated that Aurantiopileus dolosus and A. pendens have “the same kind of spores, hyphae 1a Pileus orange, basidiospores ellipsoid to and extrahymenial setae [cystidia],” but differ in broadly ellipsoid, (4.6-) 5.0-6.0 x (3.4-) the form of the fruit-body, the strigose pileus and 3.6-4.2 (-4.6) μm…....A. mayanensis the pink tubes with agglutinated tramal hyphae. 1b Pileus white, basidiospores broadly The type specimens of A. dolosus from Malaya ellipsoid to subglobose, 3.5-4.8 x 3-4 and A. pendens from Borneo both have the same μm…………………………………………………2 basidiomata form (i.e., podoporoid (dorsally substipitate)) that Corner (1989) described only 2a Tubes pale pink ………………A. dolosus in A. pendens. The reflexed habit of A. dolosus 2b Tubes white ….…………….…A. pendens reported by Corner is not obvious in the dried collection. Both have fine, radiating ridges on the Molecular considerations: While DNA- pileus surface but in A. dolosus some terminate based data from A. dolosus and A. pendens would as projecting spines or extend beyond the margin be desirable to confirm the coherence of to give a strigose appearance that is lacking in A. Aurantiopileus, molecular data are not currently pendens. available for either of these species. The age of the type collections of A. dolosus and A. pendens Both A. dolosus and A. pendens were described and the way in which they were stored make it by Corner as primarily white but in A. dolosus the likely that DNA-based data will be difficult to pileus, tubes and pores were “drab white with a obtain from these specimens, although DNA pinkish tinge;” the coloration was attributed to extraction has not yet been attempted. Further the encrustations on the cystidia. The cystidia of collecting in tropical Asia may turn up additional A. pendens “appear to have been colourless in the specimens of these species, thus affording an living state but, in material preserved in alcohol- opportunity to obtain DNA sequence data. formalin, the exudate was turned claret red” and the encrustations “turning pale pinkish ... in At the current time, all of our molecular data are potash.” Thus the pink/red tints occur in both A. based on A. mayanensis, the type of the genus. dolosus and A. pendens but under different These data (Fig. 6) place A. mayanensis firmly in circumstances. All the dried basidiomata of A. the phlebioid clade within the polyporoid clade Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10 7

Figure 6. Inferred phylogeny based on nLSU. The tree shown is one of 58 equally parsimonious trees (Length = 461, CI = 0.60, RI = 0.49). Statistical support for nodes is based on parsimony and Bayesian analyses. The asterisk after a name indicates that it is the type species of that genus. 8 Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10

(sensu Binder et al. 2005). The nLSU analysis deal of additional morphological and molecular places Aurantiopileus outside of the “core” clade work will be needed to resolve relationships of Phlebia species, one of the genera to which within the phlebioid clade, a diverse and under- Aurantiopileus is most closely related. In sampled lineage of fungi. addition, the ITS data from A. mayanensis, which could not be easily aligned to any species in the phlebioid clade, help to confirm that A. Acknowledgements: Field-work in Belize was mayanensis is an un-described species and made possible for TJB and DLL by a grant from unallied with any existing genus. The only species the National Science Foundation, Biodiversity in this analysis with ITS regions that could be Surveys and Inventories Program to the State unambiguously aligned were Phlebia acerina, P. University of New York, College at Cortland radiata (the type of genus Phlebia), and P. rufa. (DEB-0103621) and by the National Geographic As these three species are clearly closely related, Society Grant #8240-07. Dr. D. J. Lodge, USDA we consider them the “core” of genus Phlebia. Forest Service, was TJB’s co-PI on the NSF grant Sequencing of additional DNA regions will be and worked on the NGS expedition, her necessary to determine whether taxa such as P. management of logistics, contributions to tremellosa and P. centrifuga should be included obtaining specimens, and camaraderie on these within the genus Phlebia. Many of the species expeditions was indispensible. The USDA Forest within this broader group, including Service’s Center for Forest Mycology Research Crustodontia chrysocreas, Lilaceophlebia livida, provided additional logistical support. Field-work L. subserialis, Mycoacia aurea, M. fuscoatra, on Doyle’s Delight in the southern Maya and M. uda, have been included in genus Phlebia Mountains of Belize was greatly facilitated by by some authors (e.g. Lombard et al. 1975, helicopter airlift support and base camp Nakasone 1997). However, the inability to align preparation by the British Forces of Belize. ITS sequences across these taxa, coupled with the Colonel Alan Whitelaw, the commanding officer divergence of nLSU sequences, make it untenable of the Belize Forces, and Conservation Biologist at the present time to include all of these species Sharon Matola, Director of the Belize Zoo, were in one genus. ultimately responsible for making that expedition possible. Several other individuals with Unfortunately many relationships within the government and non-governmental agencies in phlebioid clade remain unresolved. The analysis Belize are kindly acknowledged for their help and in this work and in other studies (Binder et al. guidance: Dr. L. Quiroz, Ms. N. Rosado, Mr. H. 2005, Hallenberg et al. 2008) indicates that Mai, Mr. M. Windsor, and Mr. J. Pinelo of the (type species of the genus) Conservation Division of the Belize Forestry and two Gloeoporus species (G. conchoides Department in Belmopan for collecting and Mont., the type species, has not been included in export permits, Mr. Celso Poot the Station published molecular phylogenic studies) are Manager of the Tropical Education Center. The within the phlebioid clade but not closely related Doyle’s Delight expedition team of 2007 is also to Phlebia or Aurantiopileus. Other gratefully acknowledged for their help in macroscopically similar genera, such as collecting or spotting valuable specimens: Sharon Skeletocutis, have yet to be fully examined. Matola (Team Leader), Dr. D. Jean Lodge, Dr. M. (Fr.) Kotl. & Pouzar (type Catherine Aime, Mr. Jan Meerman, Mr. Gregorio species of the genus) was examined by Binder et Sho. Dr. Paul Labus, Dr. Peter Kovorik, Mr. Celso al. (2005) but could not be placed with certainty Cawich, Dr. Juan Sho, Dr. John Shuey, Ms. in any subclades of the polyporoid clade. A great Shannon Kenney, Mr. Raul Balona , Dr. Colin Ginns et al. Aurantiopileus mayanensis. Pacific Northwest Fungi 5(4): 1-10 9

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